Effects of annual grass senescence on NH4 and N-glycine uptake by blue oak (Quercus douglasii) seedlings and soil microorganisms in California oak woodland

Annual grasses are stronger competitors for available soil N than blue oak seedlings and soil microorganisms. However, little is known about the dynamics of N competition during annual grass senescence. We conducted a field experiment in a California oak woodland to study effects of annual grass senescence on N uptake by grasses, blue oak seedlings and soil microorganisms. Labeled N was applied at the beginning of April, May and of June in the form of ¹⁵NH₄⁺ or ¹⁵N-glycine. Plants and soils were harvested after 5 days (¹⁵NH₄⁺ and ¹⁵N-glycine treatments) and after 26 days (¹⁵NH₄⁺ treatment only). We evaluated effects of N form, season and labeling period on N competition among oak seedlings, annual grasses and soil microorganisms. N forms did not affect competition among grasses, oak seedlings and soil microorganisms, but more ¹⁵N was incorporated into the soil organic N pool in the ¹⁵N-glycine treatments than in the ¹⁵NH₄⁺ treatments. There were no seasonal (May vs June) effects on ¹⁵N recovery in blue oak seedlings and soil microorganisms. Plant samples from April harvest were lost. In June, when grasses were senescing, more ¹⁵N was found in the soil inorganic pool than in May. Extremely dry soils in June may have limited inorganic N availability to oak seedlings and soil microorganisms. After 26-day labeling period, ¹⁵N recovery in blue oak seedlings and the soil organic N pool significantly increased, while ¹⁵N recovery in both the soil microbial and inorganic N pools decreased compared to the 5-day labeling period. Although blue oak seedling biomass changed little from early May to late June, N concentrations in oak roots increased 53%. In contrast, annual grass biomass peaked in May, and then decreased rapidly. Our results suggest that blue oak seedlings and annual grasses have different temporal competitive abilities. Blue oak seedlings appear to have a long-term strategy for N competition. Blue oaks take up N slowly but steadily, increasing N uptake from 5 to 26 days. This extended time period has a greater positive effect on N uptake than does reduced grass uptake caused by senescence.     

CO2 and inorganic N supply modify competition for N between co-occurring grass plants, tree seedlings and soil microorganisms

Plant-plant and plant-soil interactions play a key role in determining plant community structure and ecosystem function. However, the effects of global change on the interplay between co-occurring plants and soil microbes in successional communities are poorly understood. In this study, we investigated competition for nitrogen (N) between soil microorganisms, grass plants and establishing tree seedlings under factorial carbon dioxide (CO₂) and N treatments. Fraxinus excelsior seedlings were germinated in the presence or absence of grass competition (Dactylis glomerata) at low (380 μmol mol⁻¹) or high (645 μmol mol⁻¹) CO₂ and at two levels of N nutrition in a mesocosm experiment. Pulse ¹⁵N labelling was used to examine N partitioning among plant and soil compartments. Dactylis exerted a strong negative effect on Fraxinus biomass, N capture and ¹⁵N recovery irrespective of N and CO₂ treatment. In contrast, the presence of Dactylis had a positive effect on the microbial N pool. Plant and soil responses to N treatment were of a greater magnitude compared with responses to elevated CO₂, but the pattern of Fraxinus- and microbial-N pool response to N and CO₂ varied depending on grass competition treatment. Within the Dactylis competition treatment, decreases in Fraxinus biomass in response to N were not mirrored by decreases in tree seedling N content, suggesting a shift from below- to above-ground competition. In the Dactylis-sown pots, ¹⁵N recovery could be ranked Dactylis > microbial pool > Fraxinus in all N and CO₂ treatment combinations. Inequalities between Fraxinus and soil microorganisms in terms of ¹⁵N recovery were exacerbated by N addition. Contrary to expectations, elevated CO₂ did not increase plant-microbe competition. Nevertheless, microbial ¹⁵N recovery showed a small positive increase in the high CO₂ treatment. Overall, elevated CO₂ and N supply did not interact on plant/soil N partitioning. Our data suggest that the competitive balance between establishing tree seedlings and grass plants in an undisturbed sward is relatively insensitive to CO₂ or N-induced modifications in N competition between plant and soil compartments.     

Short-term uptake of <Superscript>15</Superscript>NH<Subscript>4</Subscript><Superscript>+</Superscript> into soil microbes and seedlings of pine,spruce and birch in potted soils

Short-term competition between soil microbes and seedlings of Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies (L.) Karst.) and silver birch (Betula pendula Roth) for N was assessed in a pot study using (¹⁵NH₄)₂SO₄ as a tracer. Seedlings were grown in organic and mineral soil, collected from a podsol soil; 3.18 mg (¹⁵NH₄)₂SO₄ per pot were injected into the soil, corresponding to 4 µg ¹⁵N g^-1 d.m. (dry matter) mineral soil and 17 µg ¹⁵N g^-1 d.m. organic soil. The amounts of N and ¹⁵N in the seedlings and in microbial biomass derived from fumigation-extraction were measured 48 h after addition of ¹⁵N. In the mineral soil, 19–30% of the added ¹⁵N was found in the plants and 14–20% in the microbial biomass. There were no statistically significant differences between the tree species. In the organic soil, 74% of the added ¹⁵N was recovered in the microbial biomass in birch soil, compared to 26% and 17% in pine and spruce soils, respectively. Correspondingly, about 70% of the ¹⁵N was recovered in pine and spruce seedlings, and only 23% in birch seedlings. In conclusion, plants generally competed more successfully for added ¹⁵NH₄ ⁺ than soil microbes did. An exception was birch growing in organic soil, where the greater amount of available C from birch root exudates perhaps enabled micro-organisms to utilise more N.     

The wood-decaying fungus Hygrophoropsis aurantiaca increases P availability in acid forest humus soil, while N addition hampers this effect

We evaluated the influence of the brown rot fungus Hygrophoropsis aurantiaca on P solubility in the humus layer of a podzolic forest soil. This fungus is known to exude large amounts of oxalic acid that may stimulate weathering of minerals and increase dissolution of humus, which in turn may increase P availability in the soil surrounding the fungus. Humus was inoculated using small wooden pieces colonised by the fungus. The presence of the fungus resulted in elevated concentration of PO₄⁻ in the humus solution. In a second experiment birch seedlings grown in the same humus were able to utilise the PO₄⁻ mobilised by the fungus to increase their internal P content. The factor determining this increased P uptake and the increased available P might be oxalate produced by fungus. The acid may directly dissolve P or change organic forms of P making it more susceptible to reaction with phosphatases. This fungal effect on P solubility diminished when N was added to the soil in the form of a slow release N fertilizer (methyl urea), or when a soil with a higher soil N concentration was used. We found a strong correlation between NH₄⁺ concentration and total organic carbon in the soil solution at high NH₄⁺ concentrations, suggesting the dissolution of humus as a result of the high NH₄⁺ content in the solution. This study demonstrates that the wood-decaying fungus H. aurantiaca influences nutrient turnover in forest soil, and thereby nutrient uptake by forest trees. An intensified harvest of forest products such as whole tree harvesting may decrease the active biomass of the wood decomposers and may thereby change the availability of P and the leaching of N.     

Short-term competition between crop plants and soil microbes for inorganic N fertilizer

Agricultural systems that receive high amounts of inorganic nitrogen (N) fertilizer in the form of either ammonium (NH₄⁺), nitrate (NO₃⁻) or a combination thereof are expected to differ in soil N transformation rates and fates of NH₄⁺ and NO₃⁻. Using ¹⁵N tracer techniques this study examines how crop plants and soil microbes vary in their ability to take up and compete for fertilizer N on a short time scale (hours to days). Single plants of barley (Hordeum vulgare L. cv. Morex) were grown on two agricultural soils in microcosms which received either NH₄⁺, NO₃⁻ or NH₄NO₃. Within each fertilizer treatment traces of ¹⁵NH₄⁺ and ¹⁵NO₃⁻ were added separately. During 8 days of fertilization the fate of fertilizer ¹⁵N into plants, microbial biomass and inorganic soil N pools as well as changes in gross N transformation rates were investigated. One week after fertilization 45-80% of initially applied ¹⁵N was recovered in crop plants compared to only 1-10% in soil microbes, proving that plants were the strongest competitors for fertilizer N. In terms of N uptake soil microbes out-competed plants only during the first 4 h of N application independent of soil and fertilizer N form. Within one day microbial N uptake declined substantially, probably due to carbon limitation. In both soils, plants and soil microbes took up more NO₃⁻ than NH₄⁺ independent of initially applied N form. Surprisingly, no inhibitory effect of NH₄⁺ on the uptake and assimilation of nitrate in both, plants and microbes, was observed, probably because fast nitrification rates led to a swift depletion of the ammonium pool. Compared to plant and microbial NH₄⁺ uptake rates, gross nitrification rates were 3-75-fold higher, indicating that nitrifiers were the strongest competitors for NH₄⁺ in both soils. The rapid conversion of NH₄⁺ to NO₃⁻ and preferential use of NO₃⁻ by soil microbes suggest that in agricultural systems with high inorganic N fertilizer inputs the soil microbial community could adapt to high concentrations of NO₃⁻ and shift towards enhanced reliance on NO₃⁻ for their N supply.     

Transfer of N fixed by a legume tree to the associated grass in a tropical silvopastoral system

Below-ground transfer of nitrogen (N) fixed by legume trees to associated non-N₂-fixing crops has received little attention in agroforestry, although the importance of below-ground interactions is shown in other ecosystems. We used ¹⁵N natural abundance to estimate N transfer from the legume tree Gliricidia sepium (Jacq.) Kunth ex Walp. to C₄ grass Dichanthium aristatum (Poir.) C.E. Hubb. in a silvopastoral system, where N was recycled exclusively by below-ground processes and N₂ fixation by G. sepium was the sole N input to the system. Finding a suitable reference plant, a grass without contact with tree roots or litter, was problematic because tree roots invaded adjacent grass monocrop plots and soil isotopic signature in soil below distant grass monocrops differed significantly from the agroforestry plots. Thus, we used grass cultivated under greenhouse conditions in pots filled with agroforestry soil as the reference. A model of soil ¹⁵N fractionation during N mineralization was developed for testing the reliability of that estimate. Experimental and theoretical results indicated that 9 months after greenhouse transplanting, the percentage of fixed N in the grass decreased from 35% to <1%, due to N export in cut grass and dilution of fixed N with N taken up from the soil. The effect of soil ¹⁵N fractionation on the estimate of the reference value was negligible. This indicates that potted grass is a suitable reference N transfer studies using ¹⁵N natural abundance. About one third of N in field-grown grass was of atmospheric origin in agroforestry plots and in adjacent D. aristatum grassland invaded by G. sepium roots. The concentration of fixed N was correlated with fine root density of G. sepium but not with soil isotopic signature. This suggests a direct N transfer from trees to grass, e.g. via root exudates or common mycorrhizal networks.     

Use of 15N isotope dilution for quantification of N2 fixation associated with roots of kallar grass (Leptochloa fusca (L.))

Summary Leptochloa fusca (L.) Kunth (kallar grass) has previously been found to exhibit high rates of nitrogen fixation. A series of experiments to determine the level of biological nitrogen fixation using ¹⁵N isotopic dilution were carried out in nutrient solution and saline soil. In the nutrient solution, E. coli inoculated plants were taken as non-nitrogen-fixing control. It was observed that nearly 60%–80% of the plant N was derived from atmospheric fixation. Estimations based on the N difference method gave much lower values (18%–35%). In experiments with saline soil which was initially sterilized with chloroform fumigation, a mixed culture of N₂-fixing rhizospheric isolates from kallar grass roots was inoculated and planted to kallar grass. Uninoculated treatments were regarded as controls. The soil was previously labelled with ¹⁵N by adding cellulose and (¹⁵NH₄)₂SO₄. The results of these studies showed fixation values of 6%–32% when estimated by ¹⁵N dilution, whereas by the N difference method 54% of the plant N was estimated to be derived from fixation. This discrepancy is due to the increase in root proliferation due to inoculation, which results in greater uptake of soil N. The distribution of ¹⁵N in different fractions of the soil-N indicted isotopic dilution due to bacterial fixation of atmospheric N₂.     

Nitrate removal from drained and reflooded fen soils affected by soil N transformation processes and plant uptake

Knowledge about nitrate transformation processes and how they are affected by different plants is essential in order to reduce the loss of valuable N fertiliser as well as to prevent environmental pollution due to nitrate leaching or N₂O emission after fertilisation or the reflooding of degraded fens with nitrate-containing municipal sewage. Therefore four microcosm ¹⁵N tracer experiments were performed to evaluate the effect of common wetland plants (Phalaris arundinacea, Phragmites australis) combined with different soil moisture conditions (from dry to reflooded) on nitrate turnover processes. At the end of experiment, the total formation of gaseous N compounds was calculated using the ¹⁵N balance method. In two experiments (wet and reflooded soil conditions) the N₂O and N₂ emissions were also directly determined.Our results show that in degraded fen soils, which process mainly takes place—denitrification or transformation into organic N compounds—is determined by the soil moisture conditions. Under dry soil moisture conditions (water filled pore space: 31%) up to 80% of the ¹⁵N nitrate added was transformed into organic N compounds. This transformation process is not affected by plant growth. Under reflooded conditions (water filled pore space: 100%), the total gaseous N losses were highest (77-95% of the ¹⁵N-nitrate added) and the transformation into organic N compounds was very low (1.8% of ¹⁵N nitrate added). Under almost all soil conditions plant growth reduced the N losses by 20-25% of the ¹⁵N nitrate added due to plant uptake. The N₂ emissions exceeded the N₂O emissions by a factor of 10-20 in planted soil, and as much as 30 in unplanted soil. In the treatments planted with Phragmites australis, N₂O emission was about two times higher than in the corresponding unplanted treatment. 15% of the N₂O and N₂ formed was transported via the Phragmites shoots from the soil into the atmosphere. By contrast, Phalaris arundinacea did not affect N₂O emissions and no emission via the shoots was observed.     

NH3 Volatilization,N2O Emission and Microbial Biomass Turnover from 15N-Labeled Manure Under Laboratory Conditions

On irrigated agricultural soils from semi-arid and arid regions, ammonia (NH₃) volatilization and nitrous oxide (N₂O) emission can be a considerable source of N losses. This study was designed to test the capture of ¹⁵N loss as NH₃ and N₂O from previous and recent manure application using a sandy, calcareous soil from Oman amended one or two times with ¹⁵N labeled manure to elucidate microbial turnover processes under laboratory conditions. The system allowed to detect ¹⁵N enrichments in evolved N₂O-N and NH₃-N of up to 17% and 9%, respectively, and total N, K₂SO₄ extractable N and microbial N pools from previous and recent ¹⁵N labeled manure applications of up to 7%, 8%, and 15%. One time manured soil had higher cumulative N₂O-N emissions (141 µg kg^?1) than repeatedly manured soil with 43 µg kg^?1 of which only 22% derived from recent manure application indicating a priming effect.     

Soil microbial biomass and nitrogen transformations among five tree species of the Catskill Mountains, New York, USA

We measured soil microbial biomass nitrogen (MBN), microbial uptake of ¹⁵N, potential net mineralization and net nitrification in the laboratory to determine the influence of tree species on nitrogen (N) transformations in soils of the Catskills Mountains, New York, USA. Organic horizon soils were taken from single species plots of beech (Fagus grandifolia), hemlock (Tsuga canadensis), red oak (Quercus rubra), sugar maple (Acer saccharum) and yellow birch (Betula alleghaniensis). ¹⁵NH₄Cl was added to the soils and N pools were sampled at 1, 3, 10 and 28 days to examine microbial uptake of ¹⁵N over time. Soil MBN was about 60% lower in red oak and sugar maple soils than in the other three species. Soil pools of NO₃⁻ and rates of net nitrification were significantly greater in soils associated with sugar maple than hemlock, red oak and yellow birch. With the exception of sugar maple soils, microbial recovery of ¹⁵N was significantly greater after 10 and 28 days compared to 60 min and 1 day following ¹⁵N tracer addition. Microbial ¹⁵N recovery declined significantly within sugar maple stands within the first 3 days of incubation. Soil carbon to nitrogen ratio (C:N) was lowest in sugar maple soils and highest in red oak soils. However, correlations between soil C:N and MBN or rates of net mineralization and nitrification were not significant. Soil moisture could account for 22% of the variation in MBN and 36% of the variation in net mineralization. Soil microbial transformations of N vary among tree species stands and may have consequences for forest N retention and loss.     

Estimating N2 fixation by field-grown lupins (Lupinus angustifolius L.) using soil and plant 15N enrichment

Summary Biological N₂ fixation was estimated in a field experiment following the addition of NH₄Cl or KNO₃ to unconfined microplots (1.5 m²) at 2.5 g N m^-2 (10 atom% ¹⁵N). A model of total N and ¹⁵N accumulation in lupins and decreasing ¹⁵N enrichment in the KCl-extractable soil-N pool (0–0.15 m depth) was used to estimate the proportion of N in lupins derived from biological N₂ fixation. Estimates of N₂ fixation derived from the model were compared with ¹⁵N isotope-dilution estimates obtained using canola, annual ryegrass, and wheat as nonfixing reference plants. Biomass, total N accumulation, or ¹⁵N enrichment in the lupin and reference crops did not differ whether NH _inf4 ^sup+ or NO _inf3 ^sup- was added as the labelled inorganic-N source. The decrease in soil ¹⁵N enrichment was described by first-order kinetics, whereas total N and ¹⁵N accumulation in the lupins were described by logistical equations. Using these equations, the uptake of soil N by lupins was estimated and was then used to calculate fixed N₂. Estimates of N₂ fixation derived from the model increased from 0 at 50 days after sowing to a maximum of 0.79 at 190 days after sowing. Those based on the ¹⁵N enrichment of the NO _inf3 ^sup- pool were 10% higher than those based on the mineral-N pool. ¹⁵N isotope-dilution estimates of N₂ fixation ranged from 0.37 to 0.55 at 68 days after sowing and from 0.71 to 0.77 at 190 days after sowing. Reference plant-derived values of N₂ fixation were all higher than modelled estimates during the early states of growth, but were similar to modelled estimates at physiological maturity. The use of the model to estimate N₂ derived from the atmosphere has the intrinsic advantage that the need for a non-fixing reference plant is avoided.     

Effects of nitrogen rate and nitrogen form on glycine uptake by pakchoi (Brassica chinensis L.) under sterile culture

It is well known that plants are capable of taking up intact amino acids. However, how the nitrogen (N) rates and N forms affect amino acid uptake and amino acid nutritional contribution for plant are still uncertain. Effects of the different proportions of nitrate (NO₃^?), ammonium (NH₄⁺) and ¹⁵N-labeled glycine on pakchoi seedlings glycine uptake were investigated for 21 days hydroponics under the aseptic media. Our results showed that plant biomass and glycine uptake was positively related to glycine rate. NO₃^? and NH₄⁺, the two antagonistic N forms, both significantly inhibited plant glycine uptake. Their interactions with glycine were also negatively related to glycine uptake and glycine nutritional contribution. Glycine nutritional contribution in the treatments with high glycine rate (13.4%–35.8%) was significantly higher than that with low glycine rate (2.2%–13.2%). The high nutritional contribution indicated amino acids can serve as an important N source for plant growth under the high organic and low inorganic N input ecosystem.     

Short‐Term Recovery of Ammonium‐15Nitrogen Applied to a Temperate Forest Inceptisol and Ultisol in East Tennessee,USA

Abstract

The short‐term fate and retention of ammonium (NH₄)‐¹⁵nitrogen (N) applied to two types of forest soils in east Tennessee was investigated. Four ridgetop forests, predominantly oak (Quercus spp.), were studied. Five applications of NH₄‐¹⁵N tracer were made to the forest floor at 2‐ to 4‐week intervals over a 14‐week period in 2004. Nitrogen‐15 recovery in the forest floor, fine roots (<2 mm), and the mineral soil (0–20 cm) was calculated at 6, 21, and 42 weeks after the last application. Most of the ¹⁵N was retained in the forest floor and the mineral soil, with only small amounts (≤2%) found in roots from both soil layers. Recovery of NH₄‐¹⁵N was greater in Inceptisols, which had a wider carbon (C)‐to‐N ratio than Ultisols. For both soil types, higher NH₄‐¹⁵N recoveries and long retention times (half‐lives>100 weeks) indicated the forest floor is an effective filter for atmospheric N inputs.     

Rapid uptake of N-ammonium and glycine-C, N by arbuscular and ericoid mycorrhizal plants native to a Northern California coastal pygmy forest

While it is well established that plants are able to acquire nitrogen in inorganic form, there is less information on their ability to ‘short circuit’ the N cycle, compete with microbes, and acquire nitrogen in organic form. Mycorrhizal fungi, known to enhance nutrient uptake by plants, may play a role in organic N uptake, particularly ericoid mycorrhizas. We asked the question—Can mycorrhizal fungi increase the ability of plants to take up organic N, compared to inorganic N? Here, we report on the abilities of three plant species, ericoid mycorrhizal Rhododendron macrophyllum and Vaccinium ovatum and arbuscular mycorrhizal Cupressus goveniana ssp. pigmaea, to acquire C and/or N from an organic and an inorganic N source. All three species are native to a California coastal pygmy forest growing in acidic, low-fertility, highly organic soils. In a pot study, glycine-α¹³C, ¹⁵N and ¹⁵N-ammonium were applied to pygmy forest soil for 17 or 44 h. Ericoid mycorrhizal species did not demonstrate a preference for either inorganic or organic sources of N while Cupressus acquired more NH₄-N than glycine-N. For all species, glycine-N uptake did not increase after 17 h suggesting glycine uptake and glycine immobilization occurred rapidly. Both glycine-N and glycine-C were recovered in shoots and in roots suggesting that all species acquired some N in organic form. Regression analyses of glycine-N and glycine-C recovery in root tissue indicate that much of the glycine was taken up intact and that the minimum proportion of glycine-N recovered in organic form was 85% (Cupressus) and 70% (Rhododendron). Regressions were non-significant for Vaccinium. For all species, glycine-N remained predominantly in roots while glycine-C was transferred to shoots. In contrast, NH₄-N remained in roots of ericoid plants but was transferred to shoots of arbuscular mycorrhizal Cupressus. Since net N mineralization rates in pygmy forest soils are low, our results suggest that organic N may be an important N source for plants in this temperate coniferous ecosystem regardless of mycorrhizal type. Acquisition of amino acid C by these species also may partially offset the carbon cost to plants of hosting mycorrhizal fungi.     

Why does temperature affect relative uptake rates of nitrate, ammonium and glycine: A test with Eucalyptus pauciflora

Few studies have examined how temperature affects uptake of nitrate, ammonium and amino acids from soil. This study tests the hypothesis that cool temperatures favour uptake of the amino acid glycine while warm temperatures favour uptake of inorganic forms of N such as nitrate. We used glasshouse-grown ectomycorrhizal seedlings of the sub-alpine tree species Eucalyptus pauciflora Sieber ex Spreng. Seedlings were grown in soil (humic umbrosol, from species' habitat) that was dominated by amino acids and ammonium with only small amounts of nitrate. To examine if root physiology affects temperature responses of N uptake, we measured uptake from ¹⁵N-labelled hydrosolutions containing equimolar 100 μmol L⁻¹ mixtures of ammonium, nitrate and glycine at temperatures from 5 to 35 °C. We also examined if the effect of temperature on uptake of N forms was due to plant-microbe competition by following the fate of equimolar amounts of labelled ammonium, nitrate and glycine injected into the soil at temperatures of 5 °C and 25 °C. Hydrosolution experiments showed that uptake of glycine was favoured by warm temperatures and inorganic N by cool temperatures. In contrast, when ¹⁵N was injected into soil the uptake of glycine was favoured by low temperatures and nitrate by warm temperatures. At 25 °C, glycine was 17% of the N taken up from soil and nitrate was 51%; whereas at 5 °C glycine was 30% of the N taken up from soil and nitrate was 23%. Microbes were better competitors than seedlings for all forms of N, but temperature did not affect microbial preference for the different N forms. Hence, while microbes limit N available for plant uptake, they do not seem to be the cause of the greater plant uptake of glycine at cool temperatures and nitrate at warm temperatures. Intact uptake of glycine by plants was suggested by the positive relationship between uptake of ¹³C and ¹⁵N and detection by GC-MS of intact , ¹⁵N glycine molecules in roots. In conclusion, uptake of glycine is favoured by cool temperatures and nitrate by warm temperatures, but this is apparently not a function of root physiology or competition with soil microbes.     

Rhizosphere priming of soil organic matter by bacterial groups in a grassland soil

Plants often impact the rate of native soil organic matter turnover through root interactions with soil organisms; however the role of root-microbial interactions in mediation of the “priming effect” is not well understood. We examined the effects of living plant roots and N fertilization on belowground C dynamics in a California annual grassland soil (Haploxeralf) during a two-year greenhouse study. The fate of ¹³C-labeled belowground C (roots and organic matter) was followed under planted (Avena barbata) and unplanted conditions, and with and without supplemental N (20 kg N ha⁻¹ season⁻¹) over two periods of plant growth, each followed by a dry, fallow period of 120 d. Turnover of belowground ¹³C SOM was followed using ¹³C-phospholipid fatty acid (PLFA) biomarkers. Living roots increased the turnover and loss of belowground ¹³C compared with unplanted soils. Planted soils had 20% less belowground ¹³C present than in unplanted soils after 2 cycles of planting and fallow. After 2 treatment cycles, unlabeled soil C was 4.8% higher in planted soils than unplanted. The addition of N to soils decreased the turnover of enriched belowground ¹³C during the first treatment season in both planted and unplanted soils, however no effect of N was observed thereafter. Our findings suggest that A. barbata may increase soil C levels over time because root and exudate C inputs are significant, but that increase will be moderated by an overall faster C mineralization rate of belowground C. N addition may slow soil C losses; however, the effect was minor and transient in this system. The labeled root-derived ¹³C was initially recovered in gram negative (highest enrichment), gram positive, and fungal biomarkers. With successive growing seasons, the labeled C in the gram negative and fungal markers declined, while gram positive markers continued to accumulate labeled belowground C. The rhizosphere of A. barbata shifted the microbial community composition, resulting in greater abundances of gram negative markers and lower abundances of gram positive, actinobacteria and cyclopropyl PLFA markers compared to unplanted soil. However, the longer-term utilization of labeled belowground C by gram positive bacteria was enhanced in the rhizosphere microbial community compared with unplanted soils. We suggest that the activities of gram positive bacteria may be major controllers of multi-year rhizosphere-related priming of SOM decomposition.     

Effects of different forms of nitrogen supply on the gas exchange of young pedunculate oaks (Quercus robur L.)

The effects of different forms and concentrations of N in the rooting medium on the CO₂/H₂O gas exchange of leaves of the pedunculate oak (Quercus robur L.) were investigated. Two-year-old seedlings were grown in nutrient solutions containing low (1.8 mM) or high (4.8 mM) concentrations of NH₄⁺, 3.6 mM NO₃^?, or both NH₄⁺ and NO₃^? (1.8 mM + 1.8 mM). In various sets of plants subjected to these N treatments, the following parameters were determined: biomasses of leaves and fine roots, leaf area-related net photosynthesis at light saturation (A) and leaf conductance (g), foliar concentrations of chlorophylls, N, Ca²⁺, Mg²⁺ and K⁺ and the ash alkalinity of the leaves (as a measure of the carboxylate content). In all treatments, the leaves were equally well supplied with nutrients. Oaks grown in high NH₄⁺ concentrations produced significantly smaller leaf and root biomasses. Compared to oaks cultivated with both N forms or with low NH₄⁺ concentration, oaks grown with high NH₄⁺ supply showed lower values of A and g, but no significant differences in ash alkalinity and leaf area-related chlorophyll concentrations. Oaks fed with NO₃^? as the only N form had an intermediate biomass production, but low values of A and g. The time courses of A in the different treatments closely followed the patterns of g. In all N treatments, the same linear relationship was found between A and g, indicating that, within a rather wide range, the variation in the form and amount of supplied N does not affect the instantaneous water use efficiency of young pedunculate oaks.     

Impact of litter quality on mineralization processes in managed and abandoned pasture soils in Southern Ecuador

Tropical regions are currently undergoing remarkable rates of land use change accompanied by altered litter inputs to soil. In vast areas of Southern Ecuador forests are clear cut and converted for use as cattle pastures. Frequently these pasture sites are invaded by bracken fern, when bracken becomes dominant pasture productivity decreases and the sites are abandoned. In the present study implications of invasive bracken on soil biogeochemical properties were investigated. Soil samples (0-5 cm) were taken from an active pasture with Setaria sphacelata as predominant grass and from an abandoned pasture overgrown by bracken. Grass (C₄ plant) and bracken (C₃ plant) litter, differing in C:N ratio (33 and 77, respectively) and lignin content (Klason-lignin: 18% and 45%, respectively), were incubated in soils of their corresponding sites and vice versa for 28 days at 22 °C. Unamended microcosms containing only the respective soil or litter were taken as controls. During incubation the amount of CO₂ and its δ¹³C-signature were determined at different time intervals. Additionally, the soil microbial community structure (PLFA-analysis) as well as the concentrations of KCl-extractable C and N were monitored. The comparison between the control soils of active and abandoned pasture sites showed that the massive displacement of Setaria-grass by bracken after pasture abandonment was characterized by decreased pH values accompanied by decreased amounts of readily available organic carbon and nitrogen, a lower microbial biomass and decreased activity as well as a higher relative abundance of actinomycetes. The δ¹³C-signature of CO₂ indicated a preferential mineralization of grass-derived organic carbon in pasture control soils. In soils amended with grass litter the mineralization of soil organic matter was retarded (negative priming effect) and also a preferential utilization of easily available organic substances derived from the grass litter was evident. Compared to the other treatments, the pasture soil amended with grass litter showed an opposite shift in the microbial community structure towards a lower relative abundance of fungi. After addition of bracken litter to the abandoned pasture soil a positive priming effect seemed to be supported by an N limitation at the end of incubation. This was accompanied by an increase in the ratio of Gram-positive to Gram-negative bacterial PLFA marker. The differences in litter quality between grass and bracken are important triggers of changes in soil biogeochemical and soil microbial properties after land use conversion.     

Rhizosphere activity, grass species and N availability effects on the soil C and N cycles

We examined whether grass species and soil nitrogen (N) availability could enhance Carbon (C) and N turnover during root litter decay in grassland. Three species with increasing competitiveness (Festuca ovina, Dactylis glomerata and Lolium perenne) were grown at two N fertiliser levels in an undisturbed grassland soil, in which soil organic fractions derived for the last 9 years from Lolium root litter which was ¹³C-depleted. During the subsequent experimental year, the C turnover was calculated using the respective δ¹³C values of the old and new C in the root phytomass, in two Particulate Organic Matter (POM) fractions above 200 μm and in the lightest part of the aggregated soil fraction between 50 and 200 μm. Soil N availability was monitored during the regrowth periods with ion exchange resins (IER). The C decay rates of each particle size fraction were calculated with a simple mechanistic model of C dynamics. The N mineralisation immobilisation turnover (MIT) was characterised by dilution of ¹⁵N-labelled fertiliser in the N harvestThe C:N ratio and the residence time of C in the fractions decreased with particle size. The presence of a grass rhizosphere increased the decay rate of old C. Accumulation of new C in particle size fractions increased with species competitiveness and with N supply. Species competitiveness increased C turnover in the aggregated fraction, as a result of greater accumulation of new C and faster decay of old C. Fertiliser N increased N turnover and C mineralisation in the SOM. Species competitiveness decreased soil -N exchanged with the IER and increased dissolved organic C (DOC) content. The nature of the current rhizosphere is thus an important factor driving C and N transformations of the old root litter, in relation with grass species strategy. Plant competitiveness may stimulate the C and N turnover in the more evolved SOM fractions in a similar way to the mineral N supply.     

Size, symbiotic effectiveness and genetic diversity of field pea rhizobia (Rhizobium leguminosarum bv. viciae) populations in South Australian soils

Field pea (Pisum sativum L.) is widely grown in South Australia (SA), often without inoculation with commercial rhizobia. To establish if symbiotic factors are limiting the growth of field pea we examined the size, symbiotic effectiveness and diversity of populations of field pea rhizobia (Rhizobium leguminosarum bv. viciae) that have become naturalised in South Australian soils and nodulate many pea crops. Most probable number plant infection tests on 33 soils showed that R. l. bv. viciae populations ranged from undetectable (six soils) to 32×10³ rhizobia g⁻¹ of dry soil. Twenty-four of the 33 soils contained more than 100 rhizobia g⁻¹ soil. Three of the six soils in which no R. l. bv. viciae were detected had not grown a host legume (field pea, faba bean, vetch or lentil). For soils that had grown a host legume, there was no correlation between the size of R. l. bv. viciae populations and either the time since a host legume had been grown or any measured soil factor (pH, inorganic N and organic C). In glasshouse experiments, inoculation of the field pea cultivar Parafield with the commercial Rhizobium strain SU303 resulted in a highly effective symbiosis. The SU303 treatment produced as much shoot dry weight as the mineral N treatment and more than 2.9 times the shoot dry weight of the uninoculated treatment. Twenty-two of the 33 naturalised populations of rhizobia (applied to pea plants as soil suspensions) produced prompt and abundant nodulation. These symbioses were generally effective at N₂ fixation, with shoot dry weight ranging from 98% (soil 21) down to 61% (soil 30) of the SU303 treatment, the least effective population of rhizobia still producing nearly double the growth of the uninoculated treatment. Low shoot dry weights resulting from most of the remaining soil treatments were associated with delayed or erratic nodulation caused by low numbers of rhizobia. Random amplified polymorphic DNA (RAPD) polymerase chain reaction (PCR) fingerprinting of 70 rhizobial isolates recovered from five of the 33 soils (14 isolates from each soil) showed that naturalised populations were composed of multiple (5-9) strain types. There was little evidence of strain dominance, with a single strain type occupying more than 30% of trap host nodules in only two of the five populations. Cluster analysis of RAPD PCR banding patterns showed that strain types in naturalised populations were not closely related to the current commercial inoculant strain for field pea (SU303, ≥75% dissimilarity), six previous field pea inoculant strains (≥55% dissimilarity) or a former commercial inoculant strain for faba bean (WSM1274, ≥66% dissimilarity). Two of the most closely related strain types (≤15% dissimilarity) were found at widely separate locations in SA and may have potential as commercial inoculant strains. Given the size and diversity of the naturalised pea rhizobia populations in SA soils and their relative effectiveness, it is unlikely that inoculation with a commercial strain of rhizobia will improve N₂ fixation in field pea crops, unless the number of rhizobia in the soil is very low or absent (e.g. where a legume host has not been previously grown and for three soils from western Eyre Peninsula). The general effectiveness of the pea rhizobia populations also indicates that reduced N₂ fixation is unlikely to be the major cause of the declining field pea yields observed in recent times.